Laying the foundations for a new classification of Agaonidae (Hymenoptera: Chalcidoidea), a multilocus phylogenetic approach

Abstract

A phylogeny of the Agaonidae (Chalcidoidea) in their restricted sense, pollinators of Ficus species (Moraceae), is estimated using 4182 nucleotides from six genes, obtained from 101 species representing 19 of the 20 recognized genera, and four outgroups. Data analysed by parsimony and Bayesian inference methods demonstrate that Agaonidae are monophyletic and that the previous classification is not supported. Agaonidae are partitioned into four groups: (i) Tetrapus, (ii) Ceratosolen + Kradibia, (iii) some Blastophaga + Wiebesia species, and (iv) all genera associated with monoecious figs and a few Blastophaga and Wiebesia. The latter group is subdivided into subgroups: (i) Pleistodontes, (ii) Blastophaga psenes and neocaledonian Dolichoris, (iii) some Blastophaga and Wiebesia species, and (iv) Platyscapa, all afrotropical genera and all genera associated with section Conosycea. Eleven genera were recovered as monophyletic, six were para-or polyphyletic, and two cannot be tested with our data set. Based on our phylogeny we propose a new classification for the Agaonidae. Two new subfamilies are proposed: Tetrapusiinae for the genus Tetrapus, and Kradibiinae for Ceratosolen + Kradibia. Liporrhopalum is synonymized with Kradibia and the subgenus Valisia of Blastophaga is elevated to generic rank. These changes resulted in 36 new combinations. Finally, we discuss the hypothesis of co-speciation between the pollinators and their host species by comparing the two phylogenies.

Floral volatiles, pollinator sharing and diversification in the fig–wasp mutualism: insights from Ficus natalensis , and its two wasp pollinators (South Africa)

“…The African section, Galoglychia of Ficus, provides a unique opportunity to analyse systems in which some wasps use several host species and hosts are pollinated by several wasp species [31,32]. In KwaZulu-Natal, South Africa, E. socotrensis Mayr and E. stuckenbergi Grandi have been reported to be the sole agaonid wasps colonizing F. natalensis Hochst.…”

A living fossil fig wasp (Hymenoptera, Agaoninae) developing in extant Neotropical fig species (Moraceae: Ficus, section Pharmacosycea)

“…Due to the presence of two mandibular appendages and a membranous un-carinated gular bridge, as pleisiomorphies Agaoninae characters, the author suggests that Hexapus is the most ancestral clade of all extant fig pollinator genera; as well as, the sister clade of extant Tetrapus. This idea is reinforced by the molecular analysis of Cruaud et al (2009) …”

Chromosome numbers are not fixed in Agaonidae (Hymenoptera: Chalcidoidea)

“…Subsequent proposed phylogenies (Herre et al 1996;Jiang et al 2006;Lopez-Vaamonde et al 2009;Machado et al 2001;Weiblen 2001) were based on molecular techniques, but suffered from limited taxon and gene sampling. A recent more exhaustive study by Cruaud et al (2009) has resulted in a revised phylogeny of the family. They concluded that Agaonidae can be partitioned into four groups: the monophyletic subfamilies Tetrapusiinae (containing the single genus Tetrapus) and Kradibiinae (containing Ceratosolen and Kradibia) and two paraphyletic groups "Agaoninae" and "Blastophaginae".…”

“…The taxonomic affiliations of fig wasps within the Chalcidoidea are still uncertain (Cruaud et al 2009). The family Agaonidae has generally been restricted to the pollinator fig wasps, with non-pollinating fig wasps classified in Eurytomidae, Pteromalidae and Torymidae, but Bouček (1988) Rasplus et al (1998) re-examined the Agaonidae sensu Bouček using molecular characters they found that it contained several unrelated groups and as a result the current concept of the Agaonidae is once more restricted to fig pollinators only, with Boucek's nonpollinating subfamilies re-assigned to various other chalcid families (Cruaud et al 2009).…”

“…Early work on the phylogenetic relationships of pollinator fig wasp genera was based on morphological characters, some of which are potentially labile and subject to directional selection linked to ostiole shape and other host plant characteristics (Cruaud et al 2009;van Noort and Compton 1996). Ramirez (1978) attempted to resolve Agaonidae phylogeny based on the presence of two characters, mesosternal pollen pockets and coxal corbiculae that are associated with active pollen transfer by female fig wasps.…”